Higher Processing of Visual Information: Lecture II

1. Higher Processing of Visual Information: Lecture II • --- April 4, 2007 by Mu-ming Poo • 1. Organization of Mammalian Visual Cortices • Structure of the Primary Visual Cortex • - layering, inputs, outputs, cell types • 3. RF properties of V1 neurons • a. orientation selectivity • b. simple cell and complex cell • 4. Circuitry basis of the RFs • 5. Columnar Organization • a. orientation columns • b. ocular dominance columns

2. Brodmann’s cytoarchitectonic maps of cerebral cortex Side view of left hemisphere 18 17 Inside view of right hemisphere from the midline

3. The organization of mammalian visual cortices

4. Layer 3 Anatomy of the primary visual cortex (area 17, V1, striate cortex) V1 has six layers (2 mm thick) Layer 1 composed predominantly of dendritic and axonal connections, no neuronal cell bodies. Layers 2/3 contain excitatory neurons which project to extrastriate cortical regions Layer 4 is divided into 4A, 4B, 4Ca, and 4Cb. Layers 4Ca and 4Cb are the major recipients of LGN projections. Layers 5/6 contain excitatory projection neurons which provide feedback to LGN. Blobs --Cytochrome oxidase (CO) labeling of V1 shows CO rich regions in layer 2 & 3 termed "blobs"

5. Inputs from LGN Most LGN cells terminate in L4, (M in L4Ca, P mostly in 4Cb). Collaterals (branches)of M & P terminate in L6. K cells and some P cells terminate in blobs in L2 & 3. Output from V1 L2, 3 & 4B – feedforwad to extrastriate areas L5 - feedback to pons and superior colliculus L6 - feedback to LGN Within V1 L4  L2/3  L5  L6

6. Geniculo-cortical pathways Magnocellular pathway M-type RGCs  LGN magnocellular L1 & 2  V1 L4Ca V1 L4B • L4B cells have • Orientation selectivity • Direction selectivity • Binocular sensitivity • No color sensitivity • M channel specialized for analysis of object motion

7. Parvocellular pathway P-type RGCs  LGN parvocellular L3,4,5,6  V1 L4Cb interblob in L2 & 3 • Interblob cells in L2 & 3 have • high orientation selectivity • binocular sensitivity • small receptive fields • P channel specialized for analysis • of object shape

8. Koniocellular pathway • Blue/Yellow RGCs  LGN koniocellular layers  blobs in V1 L2 & 3. Blobs also receive inputs from LGN P layers. • Blob cells have • monocular sensitivity, • color sensitivity, • concentric RF without orientation sensitivity • Blob channel specialized for color vision

9. Cortical Cell Types Pyramidal cells Pyramidal cells -- large, pyramid shaped cell bodies, spiny dendrites, project to other areas, connect to other local neurons, all excitatory. Non-pyramidal cells -- small and stellate shape (spiny stellate or smooth stellate), local interneurons (>40 types), either excitatory (spiny) or inhibitory (smooth, few spines)

10. David Hubel (left) and Torsten Wiesel

11. Fixation point Visual field left right V1 V1 Nonuniform representation of the visual field in V1 Cortical magnification in the fovea ---- The fovea has a larger cortical representation than the peripheral.

12. + + - - + + - - + + - - - - - Cortical receptive fields Simple cells RFs have elongated nonoverlapping ON & OFF subregions and are tuned to orientation.

13. Cortical receptive fields + - - + + - - + - Complex cells RFs have overlapping ON and OFF subregions and are also tuned to orientation. Many simple and complex cells are binocular (~85%) and only respond to movement in one direction. Hypercomplex or end-stopped cells

14. Simple cell and complex cell simple cell complex cell position sensitive (no) (yes) increase response increase response decrease response (yes) length summation (yes) little response little response strong response strong response width summation (no) (yes) little or no response Little or no response orientation selectivity (yes) (yes) strong response strong response weak response weak response Little or no response Little or no response

15. Circuitry basis of V1 simple cell recetive fields + + + cortical simple cell LGN cells circuitry Hubel & Wiesel, 1962 receptive field 1. Simple cell is built up from many LGN cells 2. These LGN cells have the same center/surround structure 3. The centers of these LGN cells are distributed along a line * you can also add a set of OFF-centered LGN cells, with their centers along the OFF subregion LGN cells cortical simple cell

16. circuitry + + + complex cell complex cell simple cells simple cells Circuitry basis of V1 complex cell recetive fields receptive field 1. Complex cell is built up from many simple cells 2. These simple cells have the same preferred orientation 3. These simple cells have overlapping RFs 4. These simple cells have different arrangement of subregions

17. Columnar Organization --Cells in the same column have similar properties (RF position, orientiation preference, ocular dominance) Orientation columns Olique penetration in V1 --preferred orientation gradually shifts Vertical penetration in V1 --same preferred oritentation Ocular dominance columns Olique penetration in V1 --eye dominance shift in alternating manner Vertical penetration in V1 --same eye dominance A complete set of orientation columns is about 1 mm wide.

18. Ocular dominance columns • Monocular labeling show zebra stripes in layer IV (0.5mm wide) Cell number Ocular dominance

19. The pinwheel-like orientation maps revealed by optical imaging (Blasdel & Grinvald, 1980s). Optical imaging visualizes the changes of intrinsic optical properties of neural tissues due to neuronal activity. Iso-orientation maps of cat V1 Orientation & direction maps of monkey V1 Orientation preference map